Acclimation of Lolium temulentum to enhanced carbon dioxide concentration

Acclimation of single plants of Lolium temulentum to changing[CO₂] was studied on plants grown in controlled environmentsat 20°C with an 8 h photoperiod. In the first experimentplants were grown at 135 µ;mol m^–2 s^–1 photosyntheticphoton flux density (PPFD) at 415µl l^–1 or 550µll^–1 [CO₂] with some plants transferred from the lowerto the higher [CO₂] at emergence of leaf 4. In the second experimentplants were grown at 135 and 500 µmol m^–2 s^–1PPFD at 345 and 575 µl l^–1 [CO₂]. High [CO₂] during growth had little effect on stomatal density,total soluble proteins, chlorophyll a content, amount of Rubiscoor cytochrome f. However, increasing [CO₂] during measurementincreased photosynthetic rates, particularly in high light.Plants grown in the higher [CO₂] had greater leaf extension,leaf and plant growth rates in low but not in high light. Theresults are discussed in relation to the limitation of growthby sink capacity and the modifications in the plant which allowthe storage of extra assimilates at high [CO₂]. Key words: Lolium, carbon dioxide, photosynthesis, growth, stomatal density     

Effect of Elevated CO2 on Stomatal Density and Distribution in a C4 Grass and a C3 Forb under Field Conditions

Two common tallgrass prairie species, Andropogon gerardii, thedominant C₄ grass in this North American grassland, and Salviapitcheri, a C₃ forb, were exposed to ambient and elevated (twiceambient) CO₂ within open-top chambers throughout the 1993 growingseason. After full canopy development, stomatal density on abaxialand adaxial surfaces, guard cell length and specific leaf mass(SLM; mg cm^-2) were determined for plants in the chambers aswell as in adjacent unchambered plots. Record high rainfallamounts during the 1993 growing season minimized water stressin these plants (leaf xylem pressure potential was usually >-1·5 MPa in A. gerardii) and also minimized differencesin water status among treatments. In A. gerardii, stomatal densitywas significantly higher (190 ± 7 mm^-2; mean ±s.e.) in plants grown outside of the chambers compared to plantsthat developed inside the ambient CO₂ chambers (161 ±5 mm^-2). Thus, there was a significant 'chamber effect' on stomataldensity. At elevated levels of CO₂, stomatal density was evenlower (P < 0·05; 121 ± 5 mm^-2). Most stomatawere on abaxial leaf surfaces in this grass, but the ratio ofadaxial to abaxial stomatal density was greater at elevatedlevels of CO₂. In S. pitcheri, stomatal density was also significantlylower when plants were grown in the open-top chambers (235 ±10 mm^-2 outside vs. 140 ± 6 mm^-2 in the ambient CO₂ chamber).However, stomatal density was greater at elevated CO₂ (218 ±12 mm^-2) compared to plants from the ambient CO₂ chamber. Theratio of stomata on adaxial vs. abaxial surfaces did not varysignificantly in this herb. Guard cell lengths were not significantlyaffected by growth in the chambers or by elevated CO₂ for eitherspecies. Growth within the chambers resulted in lower SLM inS. pitcheri, but CO₂ concentration had no effect. In A. gerardii,SLM was lower at elevated CO₂. These results indicate that stomataland leaf responses to elevated CO₂ are species specific, andreinforce the need to assess chamber effects along with treatmenteffects (CO₂) when using open-top chambers.Copyright 1994, 1999Academic Press Andropogon gerardii, elevated CO₂, Salvia pitcheri, stomatal density, tallgrass prairie     

Gas Exchange and Carbohydrate and Nitrogen Concentrations in Leaves of Pascopyrum smithii (C3) and Bouteloua gracilis (C4) at Different Carbon Dioxide Concentrations and Temperatures

Pascopyrum smithii (C₃) andBouteloua gracilis (C₄) are importantforage grasses native to the Colorado shortgrass steppe. Thisstudy investigated photosynthetic responses of these grassesto long-term CO₂enrichment and temperature in relation to leafnonstructural carbohydrate (TNC) and [N]. Glasshouse-grown seedlingswere transferred to growth chambers and grown for 49 d at twoCO₂concentrations (380 and 750 µmol mol^-1) at 20 and 35°C, and two additional temperatures (25 and 30 °C) at750 µmol mol^-1CO₂. Leaf CO₂exchange rate (CER) was measuredat a plant's respective growth temperature and at two CO₂concentrationsof approx. 380 and 700 µmol mol^-1. Long-term CO₂enrichmentstimulated CER in both species, although the response was greaterin the C₃,P. smithii . Doubling the [CO₂] from 380 to 750 µmolmol^-1stimulated CER ofP. smithii slightly more in plants grownand measured at 30 °C compared to plants grown at 20, 25or 35 °C. CO₂-enriched plants sometimes exhibited lowerCER when compared to ambient-grown controls measured at thesame [CO₂], indicating photosynthetic acclimation to CO₂growthregime. InP. smithii , such reductions in CER were associatedwith increases in TNC and specific leaf mass, reductions inleaf [N] and, in one instance, a reduction in leaf conductancecompared to controls. InB. gracilis , photosynthetic acclimationwas observed more often, but significant changes in leaf metabolitelevels from growth at different [CO₂] were generally less evident.Temperatures considered optimal for growth (C₃: 20 °C; C₄:35 °C) sometimes led to CO₂-induced accumulations of TNCin both species, with starch accumulating in the leaves of bothspecies, and fructans accumulating only inP. smithii. Photosynthesisof both species is likely to be enhanced in future CO₂-enrichedand warmer environments, although responses will sometimes beattenuated by acclimation. Acclimation; blue grama (Bouteloua gracilis (H.B.K.) Lag ex Steud.); leaf nitrogen concentration; nonstructural carbohydrates; photosynthesis; western wheatgrass (Pascopyrum smithii (Rydb.) Love)     

Influence of Elevated CO2 and Temperature on the Photosynthesis and Respiration of White Clover Dependent on N2 Fixation

Single clonal plants of white clover (Trifolium repens L) grownfrom explants in a Perlite rooting medium, and dependent fornitrogen on N₂ fixation in root nodules, were grown for severalweeks in controlled environments which provided two regimesof CO₂, and temperature 23/18 °C day/night temperaturesat 680 µmol mol^–1 CO₂, (C680), and 20/15 °Cday/night temperatures at 340 µmol mol^–1 CO₂ (C340)After 3–4 weeks of growth, when the plants were acclimatedto the environmental regimes, leaf and whole-plant photosynthesisand respiration were measured using conventional infra-red gasanalysis techniques Elevated CO₂ and temperature increased ratesof photosynthesis of young, fully expanded leaves at the growthirradiance by 17–29%, despite decreased stomatal conductancesand transpiration rates Water use efficiency (mol CO₂ mol H₂O^–1)was also significantly increased Plants acclimated to elevatedCO₂, and temperature exhibited rates of leaf photosynthesisvery similar to those of C340 leaves ‘instantaneously’exposed to the C680 regime However, leaves developed in theC680 regime photosynthesised less rapidly than C340 leaves whenboth were exposed to a normal CO₂, and temperature environmentIn measurements where irradiance was varied, the enhancementof photosynthesis in elevated CO₂ at 23 °C increased graduallyfrom approx 10 % at 100 µmol m^–1 s^–1 to >27 % at 1170 µmol m^–2 s^–1 In parallel, wateruse efficiency increased by 20–40 % at 315 µmolm^–2 s^–1 In parallel, water use efficiency increasedby 20–40 % at 315 µmol m^–2 s^–1 In parallel,water use efficiency increased by 20–40 % at 315 µmolm^–2 s^–1 In parallel, water use efficiency increasedby 20–40 % at 315 µmol m^–2 s^–1 to approx100 % at the highest irradiance Elevated CO₂, and temperatureincreased whole-plant photosynthesis by > 40 %, when expressedin terms of shoot surface area or shoot weight No effects ofelevated CO₂ and temperature on rate of tissue respiration,either during growth or measurement, were established for singleleaves or for whole plants Dependence on N₂, fixation in rootnodules appeared to have no detrimental effect on photosyntheticperformance in elevated CO₂, and temperature Trifolium repens, white clover, photosynthesis, respiration, elevated CO₂, elevated temperature, water use efficiency, N₂ fixation     

Maintenance and Growth Components of Carbon Dioxide Efflux from Growing Pea Fruits

Carbon dioxide efflux from 5- to 20-day-old pea fruits was measuredfor plants grown in controlled environment at 15 °C and600 µmol s^–1 m^–2 photon flux density in a16 h photoperiod. The rate of CO₂ output per fruit increasedquickly from 0.005 to 0.018 mg CO₂ min^–1 during fruitelongation and subsequently more slowly to 0.030 mg CO₂ min^–1as the fruits inflated. On a d. wt basis the rate was highest,0.175 mg CO₂ g^–1 min^–1, in the youngest fruits anddeclined curvilinearly with increasing fruit weight to 0.02mg CO₂ g^–1 min^–1. Separation of maintenance andgrowth components was achieved by starvation methods and bymultiple regression analysis. From the latter method estimatesof the maintenance coefficient declined hyperbolically from150±8.7 mg carbohydrate g^–1 d. wt day^–1 inthe very young fruits (0.05 g) to 10.4±0.36 mg carbohydrateg^–1 d. wt day^–1 in older fruits (2.0 g). On a nitrogenbasis maintenance costs decreased from 2240 to 310 mg carbohydrateg^–1 nitrogen day^–1 while nitrogen concentrationfell from 6.7 to 3 per cent d. wt. A simple linear relationshipbetween maintenance cost per unit d. wt and nitrogen concentrationwas not observed. A growth coefficient of 50±6.7 mg carbohydrate g^–1growth (equivalent to a conversion efficiency, Y_G, of 0.95)was estimated for all fruits examined. The overall efficiency, Y, increased from a mean of 0.70 to0.85 during fruit elongation and subsequently declined to 0.80.For a given fruit weight, efficiency increased asymptoticallywith relative growth rate; both asymptote and slope of the relationshipincreased as the fruits grew. Pisum sativum L., garden pea, legume fruit, carbon dioxide efflux, maintenance respiration, growth respiration     

Response of Soybean Canopy Photosynthesis to CO2 Concentration, Light, and Temperature

Photosynthetic rates of outdoor-grown soybean (Glycine max L.Merr. cv. Bragg) canopies increased with increasing CO₂ concentrationduring growth, before and after canopy closure (complete lightinterception), when measured over a wide range of solar irradiancevalues. Total canopy leaf area was greater as the CO₂ concentrationduring growth was increased from 160 to 990 mm³ dm^–3.Photosynthetic rates of canopies grown at 330 and 660 mm³ CO₂dm^–3 were similar when measured at the same CO₂ concentrationsand high irradiance. There was no difference in ribulose bisphosphatecarboxylase/oxygenase (rubisco) activity or ribulose 1,5-bisphosphate(RuBP) concentration between plants grown at the two CO₂ concentrations.However, photosynthetic rates averaged 87% greater for the canopiesgrown and measured at 660 mm³ CO₂ dm^–3. A 10°C differencein air temperature during growth resulted in only a 4°Cleaf temperature difference, which was insufficient to changethe photosynthetic rate or rubisco activity in canopies grownand measured at either 330 or 660 mm³ CO₂ dm^–3. RuBP concentrationsdecreased as air temperature during growth was increased atboth CO₂ concentrations. These data indicate that the increasedphotosynthetic rates of soybean canopies at elevated CO₂ aredue to several factors, including: more rapid development ofthe leaf area index; a reduction in substrate CO₂ limitation;and no downward acclimation in photosynthetic capacity, as occurin some other species. Key words: CO₂ concentration, soybean, canopy photosynthesis     

Photosynthesis of White Clover Leaves as Influenced by Canopy Position, Leaf Age, and Temperature

Microswards of white clover (Trifolium repens L.) were grownin controlled environments at 10/7, 18/13 and 26/21 °C day/nighttemperatures. The vertical distribution of leaves of differentages and their rates of ¹⁴CO₂-uptake in situ were studied. Extending petioles carried the laminae of young leaves throughthe existing foliage. A final position was reached within 1/4to 1/3 of the time between unfolding and death. Newly unfoldedleaves had higher rates of ¹⁴CO₂-uptake per leaf area than olderones at the same height in the canopy. At higher temperatures,the decrease with age was faster. However, the light-photosynthesisresponse of leaves which were removed from different heightsin the canopy varied much less with leaf age than did the ratesof ¹⁴CO₂-uptake in situ. The comparison of the rates of ¹⁴CO₂-uptake in situ with thelight-photosynthesis response curves suggests that young leavesreceive more light than older ones at the same height in thecanopy. This would imply that young white clover leaves havethe ability to reach canopy positions having a favourable lightenvironment. This ability may improve the chances of survivalof white clover in competition with other species. Trifolium repens L., white clover, photosynthesis, canopy, leaf age, ¹⁴CO₂-uptake, ecotypes, temperature     

Ecophysiological Mechanisms used by Aster kantoensis, an Endangered Species, to Withstand High Light and Heat Stresses of its Gravelly Floodplain Habitat

Aster kantoensis Kitam., an endangered plant species of thefamily Compositae, is a local endemic to the gravelly floodplainsof a few rivers in central Japan. The successful growth of A.kantoensis is mainly restricted to sparsely vegetated siteswhere, due to lack of continuous vegetation, high radiant energyinput results in stressful conditions with excessive light andheat. To reveal the ecophysiological characteristics which enablethe species to cope with such environmental stresses, we measuredleaf temperature, shoot architecture and photosynthetic andtranspirational responses together with the microclimate ofthe natural habitat. Even under sunny summer conditions, theleaf temperature of A. kantoensis was much lower (35–39°C)than the soil surface temperature (max. 60°C). The relationshipbetween leaf position (height from the ground) and leaf temperatureshowed that the caulescent rosette form of A. kantoensis helpsavoid leaf overheating. Moreover, in situ gas exchange measurementsrevealed that the high transpirational capacity (as high as10 mmol H₂O m^-2s^-1) was effective in controlling leaf temperature,as long as the soil water supply was not severely limited. Sinceit has effective mechanisms to avoid the multiple stresses indigenousto its gravelly floodplain habitat, A. kantoensis can maintaina high photosynthetic rate (up to 30 µmol CO₂m^-2s^-1) withoutany midday depression under sunny summer conditions. Copyright2000 Annals of Botany Company Aster kantoensis Kitam., gravelly floodplain, high light stress, leaf temperature, photosynthesis, shoot architecture, transpiration     

Are Mistletoes Shade Plants? CO2Assimilation and Chlorophyll Fluorescence of Temperate Mistletoes and their Hosts

Mistletoes usually have slower rates of photosynthesis thantheir hosts. This study examines CO₂assimilation, chlorophyllfluorescence and the chlorophyll content of temperate host–parasitepairs (nine hosts parasitized by Ileostylus micranthus and Carpodetusserratus parasitized by Tupeia antarctica). The hosts of I.micranthus had higher mean annual CO₂assimilation (3.59 ±0.41 µmol m^-2 s^-1) than I. micranthus(2.42 ± 0.20µmol m^-2 s^-1), and C. serratus(2.41 ± 0.43 µmolm^-2 s^-1) showed higher CO₂assimilation than T. antarctica(0.67± 0.64 µmol m^-2 s^-1). Hosts saturated at significantlyhigher electron transport rates (ETR) and light levels thanmistletoes. The positive relationship between CO₂assimilationand electron transport suggests that the lower CO₂assimilationrates in mistletoes are a consequence of lower electron transportrates. When photosynthetic rates, ETR and chlorophyll a /b ratioswere adjusted for photosynthetically active radiation, hostsdid not have significantly higher CO₂assimilation (3.21 ±0.37 µmol m^-2 s^-1) than mistletoes (2.54 ± 0.41µmol m^-2 s^-1), but still had significantly higher ETRand chlorophyll a / b ratios. The electron transport rates,saturating light and chlorophyll a / b ratios of sun leavesfrom mistletoes were similar to host shade leaves. These responsesindicate that in comparison with their hosts, mistletoe leaveshave the photosynthetic characteristics of the leaves of shadeplants. Copyright 2000 Annals of Botany Company CO₂assimilation, photosynthetic active radiation (PAR), chlorophyll fluorescence, electron transport rate (ETR), photochemical quenching (q_p), non-photochemical quenching (q_n), sun and shade leaves, chlorophyll content, Ileostylus micranthus, Tupeia antarctica, New Zealand     

Photosynthesis of Stands of Tomato, Cucumber and Sweet Pepper Measured in Greenhouses under Various CO2-concentrations

The rate of net photosynthesis (P) of whole plant stands oftomato (Lycopersicon esculentum Mill.), cucumber (Cucumis sativusL.) and sweet pepper (Capsicum annuum L.) was measured in sixlong-term experiments in large greenhouses under normal operatingconditions and CO₂-concentrations between 200 and 1200 µmolmol^-1. The objective was to quantify the responses to lightand carbon dioxide and to obtain data sets for testing simulationmodels. The method of measuring canopy photosynthesis involvedan accurate estimation of the greenhouse CO₂ balance, usingnitrous oxide (N₂O) as tracer gas to determine, on-line, theexchange rate between greenhouse and outside air. The estimatedrelative error in the observed P was about ± 10%, exceptthat higher relative errors could occur under particular conditions. A regression equation relating P to the photosynthetically activeradiation, the CO₂ concentration and the leaf area index explained83-91% of the variance. The main canopy photosynthesis characteristicscalculated with the fitted regression equations were: canopyP_max 5-9 g m^-2 h^-1 CO₂ uptake; ratio P_max/LAI 1·5-3 gm^-2 h^-1; light compensation point 32-86 µmol s^-1 m^-2;light use efficiency (quantum yield) at low light 0·06-0·10µmol µmol^-1 and CO₂ compensation point 18-54 µmolmol^-1. The results were related to the prevailing conditions.Copyright1994, 1999 Academic Press Canopy photosynthesis, Capsicum annuum L., carbon dioxide, CO₂, CO₂ balance, CO₂ use efficiency, cucumber, _{Cucumis sativus} L., glasshouse, greenhouse, light use efficiency, Lycopersicon esculentum Mill., sweet pepper, tomato, tracer gas     

Some Properties of Ribulose Bisphosphate Carboxylase Extracted from Tomato Leaves

Ribulose bisphosphate carboxylase (EC 4.1.1.39 [EC] ) activity wasvery low in tomato leaf extracts unless prepared in the presenceof Mg²⁺, and polyclar AT. With young leaves, but not with fully-expanded leaves, the RuBP carboxylase activityextracted was increased by prolonged illumination of the leaves(2 h). The main effect of the light treatment was to increasethe specific activity of the enzyme but there was also a smallincrease in RuBP carboxylase protein. Tomato leaf RuBP carboxylasein extracts had specific activities in the range 0.2–0–6µmol CO₂ min^–1 mg^–-1 total protein extracted,or 0.5–1.2 µmol CO₂ min^–1 mg^–1 RuBPcarboxylase, and an apparent K_m (CO₂) at 20 ?C of 9.3 ? 1.2µM (using a of 6.407). Key words: Tomato leaf, RuBP carboxylase, Properties     

Effects of Elevated CO2 and Simulated Seasonal Changes in Temperature on the Species Composition and Growth Rates of Pasture Turves

Large turves from a ryegrass/white clover based pasture wereexposed to 350 or 700 µl l^-1 CO₂ for a period of 217 din controlled environment rooms. The temperature was increasedduring the experiment from 10/4 °C day/night to 16/10 °Cand finally to 22/16 °C. The turves were cut to a heightof 2 cm at intervals and growth rates calculated from the regrowth. Growth rates over the duration of the experiment were 8% higherat elevated CO₂; the difference between CO₂ treatments beingstatistically significant only at the highest temperature. Speciescomposition of the turves at 350 µl l^-1 CO₂ showed seasonalchanges similar to those measured in the field. The effect ofCO₂ was to exaggerate the normal decline of ryegrass at warmertemperatures and increase the proportion of white clover. About30% of the total growth rate was from other species (notablyBromus hordeaceus L. and Poa trivialis L.) and this fractionwas similar between CO₂ levels. Root mass was measured at theend of the experiment and was 50% higher at elevated CO₂. The modest above-ground response to CO₂ was a result of CO₂stimulation occurring only at the higher temperature. Becauseof the CO₂ x temperature interaction, the effect of CO₂ in temperateregions will be seasonal. When this is matched with seasonalgrowth patterns of herbage species, a complex response of pasturecommunities to CO₂ is possible. In our case, white clover wasgrowing most strongly during the period of greatest CO₂ stimulationand consequently its growth was enhanced more than that of ryegrass;however, the cooler season growth of ryegrass gives it a temporalniche which is little affected by CO₂ and this may be importantfor ryegrass stability if it is an inherently poor responderto CO₂. The results indicate that for temperate species theeffects of competition at elevated CO₂ cannot be easily determinedfrom experiments conducted at a single temperature.Copyright1994, 1999 Academic Press CO₂ enrichment, seasonal growth, species composition, turves, Trifolium repens L., Lolium perenne L., climate change     

Physiology and Growth of Wheat Across a Subambient Carbon Dioxide Gradient

Two cultivars of wheat (Triticum aestivum L.), 'Yaqui 54' and'Seri M82', were grown along a gradient of daytime carbon dioxideconcentrations ([CO₂]) from near 350-200 µmol CO₂ mol^-1air in a 38 m long controlled environment chamber. Carbon dioxidefluxes and evapotranspiration were measured for stands (plantsand soil) in five consecutive 7·6-m lengths of the chamberto determined potential effects of the glacial/interglacialincrease in atmospheric [CO₂] on C₃ plants. Growth rates andleaf areas of individual plants and net assimilation per unitleaf area and daily (24-h) net CO₂ accumulation of wheat standsrose with increasing [CO₂]. Daytime net assimilation (P_D, mmolCO₂ m^-2 soil surface area) and water use efficiency of wheatstands increased and the daily total of photosynthetic photonflux density required by stands for positive CO₂ accumulation(light compensation point) declined at higher [CO₂]. Nighttimerespiration (R_N, mmol CO₂ m^-2 soil surface) of wheat, measuredat 369-397 µmol mol^-1 CO₂, apparently was not alteredby growth at different daytime [CO₂], but R_N /P_D of stands declinedlinearly as daytime [CO₂] and P_D increased. The responses ofwheat to [CO₂], if representative of other C₃ species, suggestthat the 75-100% increase in [CO₂] since glaciation and the30% increase since 1800 reduced the minimum light and waterrequirements for growth and increased the productivity of C₃plants.Copyright 1993, 1999 Academic Press Atmospheric carbon dioxide, carbon accumulation, evapotranspiration, light compensation point, net assimilation, respiration, Triticum aestivum, water use efficiency, wheat     

Carbon Metabolism in Seagrasses: I. THE UTILIZATION OF EXOGENOUS INORGANIC CARBON SPECIES IN PHOTOSYNTHESIS

Four species of seagrasses, Halophila stipulacea, Thalassodendronciliatum, Halodule uninervis, and Syringodium isoetifolium,were investigated for their ability to utilize and CO₂ as exogenous carbon sources for photosynthesis. Ratesof photosynthesis were measured as rates of O₂ evolution ina closed system in which the pH was continuously controlled.A computer program was written to calculate the concentrationsof different carbon species as a function of pH and other specifiedexperimental conditions. Bicarbonate as well as CO₂ were readily assimilated by all fourseagrass species. Saturating concentrations of , at saturating light intensities, were 0.5–1.8 mM dependingon the species. Rates of photosynthesis under such conditionswere 0.1–0.55 µmol O₂ min^–1 mg^–1 chlorophyll.At saturating CO₂ concentrations, i.e. 0.5–1.3 mM, ratesof photosynthesis were 0.22–1.4 µmol CO₂ min^–1mg^–1 chlorophyll. Photosynthetic rates in each specieswere considerably higher when CO₂ rather than was supplied at saturating concentrations. The concentration of in natural seawater was found to be saturating, and that of CO₂ insufficient forconsiderable photosynthetic rates in these plants under thegiven conditions It was thus concluded that is the major carbon source for photosynthesis in seagrasses.     

Flag Leaf Photosynthesis of Triticum aestivum and Related Diploid and Tetraploid Species

Rates of net photosynthesis of the flag leaves of 15 genotypesof wheat and related species were measured throughout theirlife, using intact leaves on plants grown in the field. At thestage when rates were maximal, they were in general highestfor the diploid species, intermediate for the tetraploidspeciesand lowest for Triticum aestivum (means of 38, 32 and 28 mgCO₂ dm^–2 h^–1 respectively). Rates were stronglynegatively correlated with leaf area, leaf width and the meanplan area per mesophyll cell and positvely correlated with stomatalfrequency and number of veins per mm of leaf width. The differencesamong species in these attributes were mainly related to ploidylevel. It was not possible to determine the relative importanceof each anatomical feature, though the changes in stomatal frequencyhad only slight effects on stomatal conductance and the observeddifferences in rates of photosynthesis were much greater thanwould be expected from those in stomatal conductance alone. There was genetic variation in rates of light dependent oxygenevolution of isolated protoplasts and intact chloroplasts butno difference attributable to ploidy. The mean rate, 91 µmolO₂ mg^–1 chlorophyll h^–1, equivalent to 3.9 mg CO₂mg^-1chlorophyll h^-1 was considerably less than the rate of photosynthesisin comparable intact leaves, which was 7.2 mg CO₂ mg^–1chlorophyll h^–1. The total above-ground dry matter yields were least for thewild diploids T. urartu and T. thauodar and the wild tetraploidT. dicoccoides, but the other wild diploids produced as muchdry matter as the hexaploids. The prospects of exploiting differences in photosynthetic ratein the breeding of higher yielding varieties are discussed. Triticum aestivum L., wheat, Aegilops spp, photosynthesis, stomatal conductance, stomatal frequency, polyploidy     

The Carbon Economy of Rubus chamaemorus L. II. Respiration

Respiratory activity and seasonal changes in carbohydrate contentof the storage organs of Rubus chamaemorus L. have been investigated.Leaf dark respiration rate increases in a non-linear mannerfrom 0·7 mg CO₂ evolved dm^–2 h^–1 at 0 °Cto 4·6 rng CO₂ evolved dm^–2 hh^–1 at 30 °C.Root and rhizome respiration rates increase from 1 µ1O₂ uptake g^–1 fresh weight h^–1 at 0.7 ° C to10 µ10, uptake g^–1 f. wt h^–1 at 20 °C.Rhizome carbohydrate reserves decline from a September peakof 33 per cent alcohol insoluble d. wt to 16 per cent in May. The circumpolar distribution of R. chamaemorus is discussedin relation to the evidence presented here and in the precedingpaper of the series.     

Effect of Elevated CO2 on the Photosynthesis, Respiration and Growth of Perennial Ryegrass

Single, seed-grown plants of ryegrass (Lolium perenne L. cv.Melle) were grown for 49 d from the early seedling stage ingrowth cabinets at a day/night temperature of 20/15 C, witha 12 h photoperiod, and a CO₂ concentration of either 340 or680µI 1^–1 CO₂. Following complete acclimation tothe environmental regimes, leaf and whole plant CO₂ effluxesand influxes were measured using infra-red gas analysis techniques.Elevated CO₂ increased rates of photosynthesis of young, fullyexpanded leaves by 35–46% and of whole plants by morethan 50%. For both leaves and whole plants acclimation to 680µI^–1 CO₂ reduced rates of photosynthesis in bothCO₂ regimes, compared with plants acclimated to 340µll^–1. There was no significant effect of CO₂ regime onrespiration rates of either leaves or whole plants, althoughleaves developed in elevated CO₂ exhibited generally lower ratesthan those developed in 340µI I^–1 CO₂. Initially the seedling plants in elevated CO₂ grew faster thantheir counterparts in 340µI I^–1 CO₂, but this effectquickly petered out and final plant weights differed by onlyc. 10%. Since the total area of expanded and unexpanded laminaewas unaffected by CO₂ regime, specific leaf area was persistently13–40% lower in elevated CO₂ while, similarly, root/shootratio was also reduced throughout the experiment. Elevated CO₂reduced tissue nitrogen contents of expanded leaves, but hadno effect on the nitrogen contents of unexpanded leaves, sheathsor roots. The lack of a pronounced effect of elevated CO₂ on plant growthwas primarily due to the fact that CO₂ concentration did notinfluence tiller (branch) numbers. In the absence of an effecton tiller numbers, any possible weight increment was restrictedto the c. 2.5 leaves of each tiller. The reason for the lackof an effect on tillering is not known. Key words: Lolium perenne, ryegrass, elevated CO₂, photosynthesis, respiration, growth, development     

In situ Effects of Elevated Atmospheric CO2on Leaf Freezing Resistance and Carbohydrates in a Native Temperate Grassland

The objectives of this study were to quantify changes in leaffreezing resistance and carbohydrate concentrations caused bylong-term (6 years) exposure to elevated CO₂(ambient: 360 µll^-1, elevated: 600 µl l^-1) in five dominant plant speciesgrowing in situ in a native temperate grassland. Across allfive species tested from three functional groups, the mean temperatureat which all leaves were damaged (T₁₀₀) significantly (P = 0.016)increased from -9.6 to -8.5 °C under elevated CO₂ , anda similar marginally significant (P = 0.079) reduction was observedfor the mean temperature that caused 50% leaf damage (T₅₀),from -6.7 to -6.0 °C. The mean temperature at which initialleaf damage was observed (T₀) was not significantly influencedby elevated CO₂ . Although concentrations of soluble sugars(+25%,P = 0.042), starch (+53%, P < 0.001), and total non-structuralcarbohydrates (TNC, +40%, P < 0.001) were significantly higherunder elevated CO₂ , leaf freezing resistance actually decreasedunder elevated CO₂ . Concentrations of soluble sugars were positivelycorrelated with freezing resistance when viewed across all fivecommunity dominants, but within any individual species, no suchrelationships were found. We also found no evidence for ouroriginal hypothesis that increased concentrations of solublesugars increase freezing resistance. Thus, future atmosphericCO₂levels may instead increase the risk of late spring freezingdamage. Furthermore, the strong differences in freezing resistanceobserved among the species, along with decreased freezing resistance,may increase the risk of losing species that have inherentlyweak freezing resistances from the plant community. Copyright2001 Annals of Botany Company CO₂enrichment, frost hardiness, sugar, starch, total non-structural carbohydrates (TNC)     

Long-term effects of a doubled atmospheric CO2 concentration on the CAM species Agave deserti

To examine the effects of a doubled atmospheric CO₂ concentrationand other aspects of global climate change on a common CAM speciesnative to the Sonoran Desert, Agave deserti was grown under370 and 750 µmol CO₂ mol^–1 air and gas exchangewas measured under various environmental conditions. Doublingthe CO₂ concentration increased daily net CO₂ uptake by 49%throughout the 17 months and decreased daily transpiration by24%, leading to a 110% increase in water-use efficiency. Underthe doubled CO₂ concentration, the activity of ribulose-1,5-bisphosphatecarboxylase/oxygenase (Rubisco) was 11% lower, phosphoenolpyruvatecarboxylase was 34% lower, and the activated:total ratio forRubisco was 25% greater than under the current CO₂ concentration.Less leaf epicuticular wax occurred on plants under the doubledCO₂ concentration, which decreased the reflectance of photosyntheticphoton flux (PPF); the chlorophyll content per unit leaf areawas also less. The enhancement of daily net CO₂ uptake by doublingthe CO₂ concentration increased when the PPF was decreased below25 mol m^–2 d^–1 when water was withheld, and whenday/night temperatures were below 17/12 C. More leaves, eachwith a greater surface area, were produced per plant under thedoubled CO₂ concentration. The combination of increased totalleaf surface area and increased daily net CO₂ uptake led toan 88% stimulation of dry mass accumulation under the doubledCO₂ concentration. A rising atmospheric CO₂ concentration, togetherwith accompanying changes in temperature, precipitation, andPPF, should increase growth and productivity of native populationsof A. deserti. Key words: Crassulacean acid metabolism, gas exchange, global climate change, Sonoran Desert     

Effects of elevated CO2 concentrations on three montane grass species: III. Source leaf metabolism and whole plant carbon partitioning

Agrostis capillaris L.⁵, Festuca vivipara L. and Poaalpina L.were grown in outdoor open-top chambers at either ambient (340 3µmol mol^–1) or elevated (6804µmol mol^–1)concentrations of atmospheric carbon dioxide (CO₂) for periodsfrom 79–189 d. Photosynthetic capacity of source leaves of plants grown atboth ambient and elevated CO₂ concentrations was measured atsaturating light and 5% CO₂. Dark respiration of leaves wasmeasured using a liquid phase oxygen electrode with the buffersolution in equilibrium with air (21% O₂, 0.034% CO₂). Photo-syntheticcapacity of P. alpina was reduced by growth at 680 µmolmol^–1 CO₂ by 105 d, and that of F. vivipara was reducedat 65 d and 189 d after CO₂ enrichment began, suggesting down-regulationor acclimation. Dark respiration of successive leaf blades ofall three species was unaltered by growth at 680 relative to340 µmol mol^–1 CO₂. In F. vivipara, leaf respirationrate was markedly lower at 189 d than at either 0 d or 65 d,irrespective of growth CO₂ concentration. There was a significantlylower total non-structural carbohydrate (TNC) concentrationin the leaf blades and leaf sheaths of A. capillaris grown at680µmol mol^–1 CO₂. TNC of roots of A. capillariswas unaltered by CO₂ treatment. TNC concentration was increasedin both leaves and sheaths of P. alpina and F. vivipara after105 d and 65 d growth, respectively. A 4-fold increase in thewater-soluble fraction (fructan) in P. alpina and in all carbohydratefractions in F. vivipara accounted for the increased TNC content. In F. vivipara the relationship between leaf photosyn-theticcapacity and leaf carbohydrate concentration was such that therewas a strong positive correlation between photosynthetic capacityand total leaf N concentration (expressed on a per unit structuraldry weight basis), and total nitrogen concentration of successivemature leaves reduced with time. Multiple regression of leafphotosynthetic capacity upon leaf nitrogen and carbohydrateconcentrations further confirmed that leaf photosynthetic capacitywas mainly determined by leaf N concentration. In P. alpina,leaf photosynthetic capacity was mainly determined by leaf CHOconcentration. Thus there is evidence for down-regulation ofphotosynthetic capacity in P. alpina resulting from increasedcarbohydrate accumulation in source leaves. Leaf dark respiration and total N concentration were positivelycorrelated in P. alpina and F. vivipara. Leaf dark respirationand soluble carbohydrate concentration of source leaves werepositively correlated in A. capillaris. Changes in source leafphotosynthetic capacity and carbohydrate concentration of plantsgrown at ambient or elevated CO₂ are discussed in relation toplant growth, nutrient relations and availability of sinks forcarbon. Key words: Elevated CO₂, Climate change, grasses, carbohydrate partitioning, photosynthesis, respiration